A new carnivorous dinosaur from the late Jurassic Solnhofen archipelago
Ursula B. Göhlich1 and Luis M. Chiappe2
1
Dept. for Geo- and Environmental Sciences, Section Paleontology, University of Munich, Richard-
Wagner-Strasse 10, D-80333 Munich, Germany, email: u.goehlich@lrz.uni-muenchen.de.
2
The Dinosaur Institute, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los
Angeles, CA 90007, USA, email: chiappe@nhm.org
Supplementary Figure 1. Location of the type locality Schamhaupten (Southern Germany, Bavaria,
Franconian Alb). (Modified from Meyer & Schmidt-Kaler, 1989).
Supplementary Figure 2. Interpretive drawing of the bow-shaped zygapophyses of the mid-caudal (C18
to C21) vertebrae of Juravenator starki. Abbreviations: C18-C21, caudal vertebrae 18th to 21st; ha, haemal
arches; ncs, neurocentral sutures; zy, zygapophyses.
Table 1. Selected measurements of the bones of Juravenator starki (JME Sch 200).
skull
scapula (left)
humerus (left)
ulna (left)
radius (left)
manus (with claws) (left)
metacarpal I (left)
metacarpal II (left)
metacarpal III (left)
phalanx I-1 (left)
phalanx II-1 (left)
phalanx II-2 (left)
phalanx III-1 (left)
phalanx III-2 (left)
phalanx III-3 (left)
claw I (left)
claw II (left/right)
claw III (left/right)
length
82 mm
42 mm
27 mm
19.5 mm
~19.3 mm
41 mm
4.5 mm
11.5 mm
9 mm
10.5 mm
8 mm
10 mm
4 mm
4.5 mm
5.5 mm
~10 mm
9 mm / 10 mm
5.5 mm / 7 mm
length
ilium (right)
femur (right)
tibia (left/right)
pes (with claws) (left)
metatarsal I (left/right)
metatarsal II (left)
metatarsal III (left)
metatarsal IV (left/right)
metatarsal V (left/right)
phalanx I-1 (left/right)
phalanx II-1 (left)
phalanx II-2 (left/right)
phalanx III-1 (left/right)
phalanx III-2 (left/right)
phalanx III-3 (left)
phalanx IV-1 (left/right)
phalanx IV-2 (left/right)
phalanx IV-3 (right)
phalanx IV-4 (left/right)
claw I (left/right)
claw II (left/right)
claw III (left/right)
claw IV (left/right)
40 mm
52 mm
58,1 mm
63 mm
4.6 mm / 4.5 mm
26.5 mm
34.0 mm
29.6 mm / 29.8 mm
8 mm / 6,8 mm
5.8 mm / 6 mm
10.4 mm
9.0 mm / 8 mm
11.9 mm / 11.5 mm
8.1 mm / 8 mm
7.4 mm
7.4 mm / 7 mm
5.5 mm / 6.5 mm
4.5 mm
4.2 mm / 4 mm
6.0 mm / 3.5 mm
10.7 mm / 11.5 mm
7.4 mm / 6.6 mm
7.2 mm / 5.8 mm
Cladistic Analysis. Morphological data of Juravenator starki was included in a matrix of 189
characters (see Appendix 1). The 35 terminal taxa included Juravenator starki, 30 coelurosaur
species (Microraptor zhaoianus, Sinornithosaurus millenii, Velociraptor mongoliensis, Shuuvuia
deserti, Mononykus olecranus, Patagonykus puertai, Confuciusornis sanctus, Archeopteryx
lithographica, Caudipteryx zoui, Conchoraptor gracilis, Citipati osmolskae, Shenzhousaurus
orientalis, Ornithomimus edmonticus, Gallimimus bullatus, Troodon formosus, Saurornithoides
mongoliensis, Sinovenator changii, Alxasaurus elesitaiensis, Erlikosaurus andrewsi,
Albertosaurus sarcophagus, Tyrannosaurus rex, Ornitholestes hermanni, Huaxiagnathus
orientalis, Sinosauropteryx prima, Compsognathus longipes, Mirischia asymmetrica, Scipionyx
samniticus, Nqwebasaurus thwazi, Coelurus fragilis, Nedcolbertia justinhoffmanni), two basal
tetanurans (Allosaurus fragilis and Sinraptor dongi), and two non-tetanuran theropods
(Coelophysis bauri and Dilophosaurus wetherilli). An initial analysis of all these taxa resulted in
a strict consensus cladogram (107 trees of 534 steps) with limited resolution at the base of the
tree. In this tree, Juravenator was nested with a number of other basal coelurosaurs in a large
polichotomy (Supplementary Figure 3). A second analysis omitting seven taxa, either less
relevant (i.e., Alxasaurus elesitaiensis, Erlikosaurus andrewsi) or largely incomplete (i.e.,
Mirischia asymmetrica, Scipionyx samniticus, Nqwebasaurus thwazi, Coelurus fragilis,
Nedcolbertia justinhoffmanni), resulted in the cladogram discussed and figured in the text (Figure
4 of text). Both analyses were conducted using 5 replications of random addition sequences of the
taxa creating Wagner trees, each followed by TBR (Tree Bisection Reconnection) branch
swapping and 100 iterations of jackknife ratchet (Nixon, 1999), collapsing the trees on TBR
rearrangements. This search strategy was implemented using Goloboff’s (1993) program NONA.
Given the size of the present data, this strategy is almost certain to produce an entirely accurate
consensus tree (see Goloboff and Farris, 2001). All cladograms used Coelophysis bauri as the
root.
Supplementary Figure 3. Strict consensus tree (534 steps) resultant from the cladistic analysis
of the data matrix listed in the Appendix 2.
Appendix 1. Character list for cladistic analysis based on Hwang et al. (2004). Of the 222 characters used
by Hwang et al. (2004), we have omitted 43 characters (i.e., Hwang et al.’s characters 6, 30, 32, 37, 45,
49, 50, 55, 92, 95, 99, 100, 103, 107, 109, 119, 122, 134, 135, 141, 143, 156, 163, 166, 168, 169, 170,
174, 177, 182, 189, 191, 198, 199, 203, 204, 210, 211, 214, 215, 216, 219, 220). We have also added 10
characters to the remaining list and modified (description and/or states) a number of Hwang et al.’s (2004)
original characters (see below). The states of multistate characters 40, 44, 70, 77, 89, 90, 110, 113, 120,
121, 148, 159, 167, 168, 172, and 186 were treated as ‘ordered’ (additive characters) (see below).
Integument
1. Vaned feathers on forelimb: symmetric (0) or asymmetric (1).
Skull
2. Orbit in lateral or dorsolateral view: round (0) or dorsoventrally elongate (1).
3. Anterior process of postorbital: projects into orbit (0) or does not project into orbit (1).
4. Frontal (anterior) process of postorbital in lateral view: straight (0) or anterodorsally curved with
concave dorsal border (1).
5. Postorbital bar: parallels quadrate, lower temporal fenestra rectangular in shape (0) or jugal and
postorbital approach or contact quadratojugal to constrict lower temporal fenestra (1).
6. Crista interfenestralis: confluent with lateral surface of prootic and opisthotic (0) or distinctly depressed
within middle ear opening (1).
7. Subotic recess (pneumatic fossa ventral to fenestra ovalis): absent (0) or present (1).
8. Basisphenoid recess: located between basisphenoid and basioccipital (0), entirely within basisphenoid
(1), or absent (2).
9. Posterior opening of basisphenoid recess: single (0) or divided into two small, circular foramina by a
thin bar of bone (1).
10. Base of cultriform process (parasphenoid rostrum): not highly pneumatized (0) or expanded and
pneumatic (parasphenoid bulla) (1).
11. Basipterygoid processes: projecting ventrally or anteroventrally (0) or lateroventrally (1).
12. Basipterygoid processes: well developed, extending as a distinct process from the base of the
basisphenoid (0) or abbreviated or absent (1).
13. Basipterygoid processes: solid (0) or hollow (1).
14. Basipterygoid recesses on dorsolateral surface of basipterygoid processes: absent (0) or present (1).
15. Depression for pneumatic recess on prootic: absent (0) or present as dorsally open fossa on
prootic/opisthotic (1) or present as deep, posterolaterally directed concavity (2).
16. Accessory tympanic recess dorsal to crista interfenestralis: absent (0), small pocket present (1), or
extensive with indirect pneumatization (2).
17. Caudal (posterior) tympanic recess: absent (0), present as opening on anterior surface of paroccipital
process (1), or extends into opisthotic posterodorsal to fenestra ovalis, confluent with this fenestra (2).
18. Exits of cranial nerves X-XII: flush with surface of exoccipital (0) or cranial nerve exits located
together in a bowl-like basisphenoid depression (1).
19. Maxillary process of premaxilla: contacts nasal to form posterior border of external nares (0),
maxillary process reduced so that maxilla participates broadly in external naris (1), or maxillary
process extends posteriorly to separate maxilla from nasal posterior to nares (2).
20. Internarial bar: rounded (0) or flat (1).
21. Premaxillary subnarial depth: main body as long or longer rostrocaudally than high dorsoventrally (0)
or very high, much shorter rostrocaudally than dorsoventrally (1). In birds, the main body of the
premaxilla is not entirely ventral to the external nares—the latter has migrated caudally in more or less
degree. For this reason, we have scored Archaeopteryx and Confuciusornis as non-comparable. (added)
22. Crenulate margin on buccal edge of premaxilla: absent (0) or present (1).
23. Caudal margin of naris: farther rostral than (0), or nearly reaching or overlapping (1), the rostral
border of the antorbital fossa.
24. Premaxillary symphysis: acute, V-shaped (0) or rounded, U-shaped (1).
25. Secondary palate: formed by premaxilla only (0) or by premaxilla, maxilla, and vomer (1).
26. Palatal shelf of maxilla: flat (0) or with midline ventral ‘tooth-like’ projection (1).
27. Pronounced, round accessory antorbital fenestra (maxillary fenestra): absent (0) or present (1).
28. Accessory antorbital fenestra (maxillary fenestra): situated at rostral border of antorbital fossa (0) or
situated posterior to rostral border of fossa (1).
29. Tertiary antorbital fenestra (promaxillary fenestra): absent (0) or present (1).
30. Maxillary rostral margin: convex (0), concave, rostral ramus equal or shorter than higher (1), concave,
rostral ramus longer than higher (2). (added)
31. Nasal participation in antorbital fossa: minimal or absent (0), present (1). (added)
32. Narial region: apneumatic or poorly pneumatized (0) or with extensive pneumatic fossae, especially
along posterodorsal rim of fossa (1).
33. Jugal tall beneath lower temporal fenestra, twice or more as tall dorsoventrally as it is wide
transversely (0) or rod-like (1). We coded the very short quadratojugal process of the jugal of
Archaeopteryx (Elzanowski and Wellnhofer, 1996) as non-comparable.
34. Jugal pneumatic recess in posteroventral corner of antorbital fossa: present (0) or absent (1).
35. Medial jugal foramen: present on medial surface ventral to postorbital bar (0) or absent (1).
36. Quadratojugal: without horizontal process posterior to ascending process (reversed “L” shape) (0) or
with process (i.e., inverted ‘T’ or ‘Y’ shape) (1).
37. Supraorbital crests on lacrimal in adult individuals: absent (0) or dorsal crest above orbit (1) or lateral
expansion anterior and dorsal to orbit (2).
38. Enlarged foramen or foramina opening laterally at the angle of the lacrimal: absent (0) or present (1).
39. Lacrimal posterodorsal process: absent (inverted ‘L’ shaped) (0) or lacrimal ‘T’ shaped in lateral view
(1) or anterodorsal process much longer than posterior process (2).
40. Prefrontal dorsal exposure: large, similar to or larger than that of lacrimal (0) or greatly reduced in size
(1) or absent (2). (ordered)
41. Anterior end of frontals: narrow, forming a wedge between nasals (0) or ending abruptly anteriorly,
suture with nasal transversely oriented (1).
42. Anterior emargination of supratemporal fossa on frontal: straight or slightly curved (0) or strongly
sinusoidal and reaching onto postorbital process (1) (see Currie 1995).
43. Postorbital process of frontal in dorsal view: smooth transition from orbital margin (0) or sharply
demarcated from orbital margin (1) (see Currie 1995).
44. Dorsal surface of parietals: flat, lateral ridge borders supratemporal fenestra (0), parietals dorsally
convex with very low sagittal crest along midline (1), or dorsally convex with well developed sagittal
crest (2). (ordered)
45. Parietals: separate (0) or fused (1).
46. Descending process of squamosal: parallel (0) or nearly perpendicular (1) to quadrate shaft.
47. Dorsal process of quadrate: single headed (0) or with two distinct heads, a lateral one contacting the
squamosal and a medial head contacting the braincase (1).
48. Quadrate: vertical (0) or strongly inclined anteroventrally so that distal end lies far forward of
proximal end (1).
49. Quadrate: solid (0) or hollow, with depression on posterior surface (1).
50. Quadrate head in lateral view: covered by squamosal (0) or exposed, quadrate cotyle of squamosal
open laterally (1).
51. Lateral border of quadrate shaft: straight in caudal view (0) or with lateral tab that touches squamosal
and quadratojugal above an enlarged quadrate foramen (1), when these bones are in contact to each
other. Although the squamosal and quadratojugal of Shuvuuia do not contact each other (Chiappe et al.,
2002), the lateral border of the quadrate of this taxon is laterally notched and not straight—thus, we
have still coded as “1” despite the absence of contact between the squamosal and quadratojugal.
(states modified)
52. Occipital condyle: without constricted neck (0) or subspherical with constricted neck (1).
53. Paroccipital process: elongate and slender, with dorsal and ventral edges nearly parallel (0) or short
and deep, with convex distal end (1).
54. Paroccipital process: straight, projects laterally or posterolaterally (0) or distal end curves ventrally,
pendant (1).
55. Dorsal edge of paroccipital process: straight (0) or twisted rostrolaterally at distal end (1) (see Currie
1995).
56. Ectopterygoid: with constricted opening into fossa (0) or with open ventral fossa in the main body of
the element (1).
57. Ventral recess on ectopterygoid: absent (0) or present (1).
58. Flange of pterygoid: well developed (0) or reduced in size or absent (1).
59. Palatine and ectopterygoid: separated by pterygoid (0) or contact each other (1) (see Currie 1995).
60. Palatine: tetraradiate, with jugal process (0) or triradiate, jugal process absent (1).
61. Suborbital fenestra: similar in length to orbit (0) or reduced in size (less than one quarter orbital
length) or absent (1).
62. Depression (possibly pneumatic) on ventral surface of postorbital process of laterosphenoid: absent (0)
or present (1).
63. Basal tubera: set far apart, level with or beyond lateral edge of occipital condyle and/or foramen
magnum (may connected by a web of bone or separated by a large notch) (0) or tubera small, directly
below condyle and foramen magnum, and separated by a narrow notch (1).
Mandible
64. Symphyseal region of dentary in occlusal view: almost straight (symphyseal surface aligned with
medial margin of mandible) (0) or medially recurved (1). (states modified; see Rauhut, 2003)
65. Dentary symphyseal region: in line with main part of buccal edge (0) or symphyseal end downturned
(1).
66. Mandibular coronoid prominence: absent (0) or present (1).
67. Posterior end of dentary: straight or only slightly concave (minimal or no development of process
dorsal to mandibular fenestra, if present) (0) or strongly forked (with strong process above mandibular
fenestra, if present) (1) (see Rauhut, 2003). (states modified)
68. Labial face of dentary: flat (0) or with lateral ridge and inset tooth row (1).
69. Shape of dentary in lateral view: subtriangular (0) or with subparallel dorsal and ventral edges (1) (see
Currie 1995).
70. Dentary: fully toothed (0) or only with teeth rostrally (1) or edentulous (2). (ordered)
71. External mandibular fenestra: large (0), reduced or absent (1). (added)
72. Nutrient foramina on external surface of dentary: superficial (0) or lie within deep groove (1).
73. Large external mandibular fenestra: oval (0) or subdivided by a spinous rostral process of the
surangular (1). (states modified)
74. Internal mandibular fenestra: absent or small and slit-like (0) or large and rounded (1) (see Currie
1995). (states modified)
75. Foramen in lateral surface of surangular rostral to mandibular articulation, absent (0) or present (1).
76. Lateral exposure of splenial: not widely exposed (0) or exposed as a broad triangle between dentary
and angular (1).
77. Coronoid ossification: large (0), or only a thin splint (1), or absent (2). (ordered)
78. Laterally inclined flange along dorsal edge of surangular for articulation with lateral process of lateral
quadrate condyle: absent (0) or present (1).
79. Angular exposed almost to end of mandible in lateral view: reaches or almost reaches articular (0) or
excluded from posterior end angular suture turns ventrally and meets ventral border of mandible rostral
to glenoid (1).
80. Articular: without elongate, slender medial, posteromedial, or mediodorsal process from retroarticular
process (0) or with process (1).
81. Retroarticular process: short, stout (0) or elongate and slender (1).
82. Mandibular articulation surface: as long as distal end of quadrate (0) or twice or more as long as
quadrate surface, allowing anteroposterior movement of mandible (1).
83. Jaws occlusion: throughout their length (0) or jaws diverge rostrally due to kink and downward
deflection in dentary buccal margin (1).
Dentition
84. Premaxilla teeth: present (0) or absent (1).
85. Premaxillary teeth: all or some serrated (0), without serrations (1). (added)
86. Second premaxillary tooth: approximately equivalent in size to other premaxillary teeth (0) or
markedly larger than third and fourth premaxillary teeth (1) (see Currie 1995).
87. Maxillary teeth: present (0) or absent (1).
88. Caudal extension of upper tooth row: to within the level of orbit (0), to rostral margin of, or anterior to
the orbit (1). (added)
89. Maxillary and dentary teeth: serrated (0) or some without serrations anteriorly (except at base in
Saurornithoides mongoliensis) (1) or all without serrations (2). (ordered)
90. Dentary and maxillary teeth: large, less than 25 in dentary (0) or moderate number of small teeth (2530 in dentary) (1) or teeth relatively small, and numerous (more than 30 in dentary) (2). (ordered)
91. Serration denticles: large (0) or small (1).
92. Serrations: simple, denticles convex (0) or distal and often mesial edges of teeth with large, hooked
denticles that point toward the tip of the crown (1).
93. Teeth constriction between root and crown: present (0) or absent (1).
94. Dentary teeth: evenly spaced (0) or anterior dentary teeth smaller, more numerous, and more closely
appressed than those in middle of tooth row (1).
95. Dentary interdental plates: absent (0) or present medially between teeth (1).
96. Premaxillary tooth crowns in cross section: sub-oval to sub-circular (0) or asymmetrical (D-shaped in
cross section) with flat lingual surface (1).
Axial skeleton
97. Axial epipophyses: absent or poorly developed, not extending past posterior rim of postzygopophyses
(0) or large and posteriorly directed, extend beyond postzygapophyses (1).
98. Axial neural spine: flared transversely (0) or compressed mediolaterally (1).
99. Anterior cervical centra: level with or shorter than posterior extent of neural arch (0) or extending
beyond posterior limit of neural arch (1).
100. Carotid process on posterior cervical vertebrae: absent (0) or present (1).
101. Cervical vertebrae pneumaticity: two pneumatic foramina arranged horizontally (0) or a single
pneumatic foramen (1). (added)
102. Anterior cervical centra: subcircular or square in anterior view (0) or distinctly wider than high,
kidney shaped (1).
103. Cervical and anterior dorsal vertebrae: amphiplatyan (0) or opisthocoelous (1).
104. Anterior dorsal vertebrae: without prominent hypapophyses (0) or with large hypapophyses (1).
105. Hyposphene-hypantrum articulations in dorsal vertebrae: absent (0) or present (1).
106. Zygapophyses of dorsal vertebrae: abutting one another above neural canal, opposite hyposphenes
meet to form lamina (0), or zygapohyses placed lateral to neural canal and separated by groove for
interspinuous ligaments, hyposphenes separated (1).
107. Presacral vertebrae pneumaticity: cervical vertebrae but not dorsal vertebrae pneumatic (0) or all
presacral vertebrae pneumatic (1).
108. Apices of neural spines of dorsal vertebrae: not expanded (0) or expanded transversally to form a
‘spine table’ (1).
109. Neural spines on posterior dorsal vertebrae in lateral view: rectangular or square (0) or
anteroposteriorly expanded distally, fan-shaped (1).
110. Number of sacral vertebrae: 4 or 5 (0), or 6 or 7 (1), or 8 or more (2). (states modified; ordered) Our
coding for Gallimimus follows Osmolska et al. (1972) and differs from that given by Hwang et al.
(2004). (states modified)
111. Sacral vertebrae: with unfused zygapophyses (0) or with fused zygapophyses forming a sinuous ridge
in dorsal view (1).
112. Ventral surface of posterior sacral centra: convex and gently rounded (0), or ventrally flattened,
sometimes with a shallow sulcus (1), or centrum strongly constricted transversely, ventral surface
keeled (2).
113. Pleurocoels on sacral vertebrae: absent (0), or present on anterior sacrals only (1), or present on all
sacrals (2). (ordered)
114. Last sacral centrum: with flat posterior articulation surface (0) or convex articulation surface (1).
115. Caudal vertebrae: with distinct transition point, from shorter centra with long transverse processes
proximally to longer centra with small or no transverse processes distally (0) or vertebrae
homogeneous in shape, without transition point (1).
116. Transition point in caudal series: begins distal to the 10th caudal (0) or at or proximal to the 10th
caudal vertebra (1).
117. Anterior caudal centra: tall, oval in cross section (0), with box-like centra in caudals I-V (1), or
anterior caudal centra laterally compressed with ventral keel (2).
118. Neural spines of caudal vertebrae: simple, undivided (0) or separated into anterior and posterior alae
throughout much of caudal sequence (1).
119. Prezygapophyses of distal caudal vertebrae: between 1/3 and whole centrum length (0), or with
extremely long extensions of the prezygapophyses (up to 10 vertebral segments long in some taxa) (1),
or strongly reduced as in Archaeopteryx lithographica (2).
120. Number of caudal vertebrae: more than 40 (0), between 25 and 40 (1), or no more than 25 (2).
(ordered)
121. Distal caudal chevrons: simple (0), bifurcate anteriorly (1), or bifurcate at both ends (2). (ordered)
122. Shaft of cervical ribs: slender and longer than vertebra to which they articulate (0) or broad and
shorter than vertebra (1).
123. Ossified uncinate processes: absent (0) or present (1).
124. Ossified ventral rib segments: absent (0) or present (1).
125. Lateral gastral segment: shorter (0) or longer (1) than medial one in each arch.
126. Ossified sternal plates: separate in adults (0) or fused (1).
127. Sternum: without distinct lateral xiphoid process posterior to costal margin (0) or with lateral xiphoid
process (1).
128. Coracoidal grooves on anterior edge of sternum: present (0) or absent (1).
129. Articular facet of coracoid on sternum (conditions may be determined by the articular facet on
coracoid in taxa without ossified sternum): anterolateral or more lateral than anterior (0); almost
anterior (1).
Pectoral Girdle
130. Hypocleidium on furcula: absent (0) or present (1).
131. Acromion margin of scapula: continuous with blade (0) or anterior edge laterally everted (1).
132. Coracoid in lateral view: subcircular, with shallow ventral blade (0) or subquadrangular with
extensive ventral blade (1) or shallow ventral blade with elongate posteroventral process (2).
133. Posterior edge of coracoid: not or shallowly indented below glenoid (0), or posterior edge of coracoid
deeply notched just ventral to glenoid, glenoid lip everted (1).
134. Scapula and coracoid form a continuous arc in posterior and anterior views (0) or coracoid inflected
medially, scapulocoracoid ‘L’ shaped in lateral view (1).
135. Glenoid fossa faces posteriorly or posterolaterally (0) or laterally (1).
136. Scapula longer (0) or shorter (1) than humerus.
Forelimb
137. Deltopectoral crest large and distinct, proximal end of humerus quadrangular in anterior view (0),
deltopectoral crest less pronounced, forming an arc rather than being quadrangular (1), deltopectoral
crest very weakly developed, proximal end of humerus with rounded edges (2), deltopectoral crest
extremely long (3), or proximal end of humerus extremely broad, triangular in anterior view (4).
138. Olecranon process: weakly developed (0) or distinct and large (1).
139. Proximal surface of ulna: a single continuous articular facet (0) or divided into two distinct fossae
separated by a median ridge (1).
140. Lateral proximal carpal (ulnare?): oval to quadrangular (0) or triangular in proximal view (1). (states
modified)
141. Distal carpals in contact with metacarpals: three carpals, one partially capping the base of
metacarpals I and II, another capping the base of metacarpal III, and a small third one capping
metacarpal IV (0); two carpals, one covering the base of metacarpal I (and perhaps contacting
metacarpal II) and the other covering the base of metacarpal II (1); or a single distal carpal capping
metacarpals I and II (2). (states modified)
142. Distal carpals: not fused to metacarpals (0) or fused to metacarpals, forming carpometacarpus (1).
143. Semilunate distal carpal: well developed, covering all of proximal ends of metacarpals I and II (0),
small, covers about half of base of metacarpals I and II (1), or covers bases of all metacarpals (2).
144. Metacarpal I approximately half or less than half the length of metacarpal II, and longer
proximodistally than wide transversely (0), distinctly more than half the length or subequal in length to
metacarpal II (1), or very short and wider transversely than long proximodistally (2). (states modified)
145. Shaft diameter of manual phalanx I-1: less (0) or greater (1) than shaft diameter of radius.
146. Fourth manual digit: present, albeit largely limited to its metacarpal (0) or complete absent (1).
(added)
147. Third manual digit: present, phalanges present (0) or reduced to no more than metacarpal splint (1).
148. Manual unguals: strongly curved, with large flexor tubercles (0), weakly curved with weak flexor
tubercles displaced distally from articular end (1), or straight with weak flexor tubercles displaced
distally from articular end (2). (ordered)
149. Unguals on all digits generally similar in size (0) or digit I bearing large ungual and unguals of other
digits distinctly smaller (1).
150. Proximodorsal ‘lip’ on some manual unguals (a transverse ridge immediately dorsal to the
articulating surface): absent (0) or present (1).
Pelvic Girdle
151. Ventral edge of anterior ala of ilium: straight or gently curved (0) or ventral edge hooked anteriorly
(1). (states modified)
152. Preacetabular part of ilium: roughly as long as postacetabular part of ilium (0) or preacetabular
portion of ilium markedly longer (more than 2/3 of total ilium length) than postacetabular part (1).
153. Supraacetabular crest on ilium: prominent, capping femoral head (0) reduced or absent (1). (states
modified).
154. Postacetabular ala of ilium in lateral view: squared (0) or acuminated (1).
155. Postacetabular blades of ilia in dorsal view: parallel (0) or diverge posteriorly (1).
156. Tuber along dorsal edge of ilium, dorsal or slightly posterior to acetabulum: absent (0) or present (1).
157. Brevis fossa: shelf-like (0) or deeply concave with lateral overhang (1).
158. Antitrochanter posterior to acetabulum: absent or poorly developed (0) or prominent (1).
159. Cuppedicus fossa: deep, ventrally concave (0) or fossa shallow or flat, with no lateral overhang (1) or
absent (2). (ordered)
160. Distal extension of pubic peduncle: similar to that of ischial peduncle (0) or much greater than that of
ischial peduncle (1). (added)
161. Posterior edge of ischium: straight (0) or with caudal (proximodorsal) process (1).
162. Hooked ventral projection of pubis at the level of its articulation with the ischium: absent (0), present
(1). (added)
163. Obturator process of ischium: absent (0), proximal in position (1), located near middle of ischiadic
shaft (2), or located at distal end of ischium (3).
164. Ischium more than two-thirds (0) or two-thirds or less of pubis length (1).
165. Distal ends of ischia form symphysis (0) or they are separated and do not form a symphysis (1).
(states modified)
166. Ischial boot (expanded distal end): present (0) or absent (1). We coded the autapomorphic condition
of Archaeopteryx (Wellnhofer, 1973) as non-comparable.
167. Pubis propubic (0), pubis vertical to subvertical (1), or pubis posteriorly oriented (opisthopubic) (2).
(states modified; ordered)
168. Pubic boot projects anteriorly and posteriorly (0), with little or no anterior process and strong
posterior process (1), or minimal or no anteroposterior projections (2). (states modified; ordered)
169. Pubic shaft straight or anteriorly convex (0) or distal end curves anteriorly, anterior surface of shaft
concave (1). (states modified)
170. Pubic apron about half of pubic shaft length (0) or less than 1/3 of shaft length (1).
Hindlimb
171. Femoral head: without fovea capitalis (for attachment of capital ligament) (0) or circular fovea
present in center of medial surface of head (1).
172. Lesser trochanter: short, ending proximally far distal to greater trochanter (0), tall and separated from
greater trochanter by deep cleft (1), tall and separated from greater trochanter by small groove (2), or
completely fused (or absent) to form crista trochanteris (3). (states modified; ordered)
173. Posterior trochanter: absent or represented only by rugose area (0) or posterior trochanter distinctly
raised from shaft, mound-like (1).
174. Fourth trochanter on femur: present (0) or minimally developed or absent (1).
175. Accessory trochanteric crest distal to lesser trochanter of femur: absent (0) or present (1).
176. Anterior surface of femur proximal to medial distal condyle: without longitudinal crest (0) or crest
present extending proximally from medial condyle on anterior surface of shaft (1).
177. Popliteal fossa on distal end of femur: open distally (0) or closed off distally by contact between
distal condyles (1).
178. Fibula reaches proximal tarsals (0) or short, tapering distally, and not in contact with proximal tarsals
(1).
179. Deep oval fossa on medial surface of fibula near proximal end: absent (0) or present (1).
180. Medial cnemial crest: absent (0) or present on proximal end of tibia (1).
181. Ascending process of the astragalus: very short, stout, and craniocaudally broad (0), tall and broad,
covering most of anterior surface of distal end of tibia (1), process short and slender, covering only
lateral half of anterior surface of tibia (2), or ascending process tall with medial notch that restricts it to
lateral side of anterior face of distal tibia (3). (states modified)
182. Ascending process of astragalus: confluent with condylar portion (0) or separated by transverse
groove or fossa across base (1).
183. Astragalus and calcaneum: separate from tibia (0) or fused to each other and to the tibia in late
ontogeny (1).
184. Distal tarsals: separate, not fused to metatarsals (0) or at least some fused to metatarsals in adults (1).
(states modified)
185. Metatarsals: not co-ossified (0) or proximally co-ossified (1). (states modified)
186. Proximal surface of metatarsal IV: subequal to II in size, proximal end of metatarsal III visible
between metatarsals II and IV in anterior view (0), metatarsal III pinched between metatarsals II and
IV, the latter two contacting one another proximally in front of III (1), or metatarsal III does not reach
proximal end of metatarsus (2). (ordered)
187. Ungual and penultimate phalanx of pedal digit II similar to those of III (0) or penultimate phalanx
highly modified for extreme hyper-extension, ungual more strongly curved and about 50% larger than
that of III (1).
188. Metatarsal I articulates on the central half of the medial surface of metatarsal II (0), attaches to
posterior surface of distal quarter of metatarsal II (1), or articulates to medial surface of metatarsal II
near its proximal end (2), or metatarsal I absent (3). (states modified)
189. Foot symmetrical (0) or asymmetrical with slender metatarsal II and very robust metatarsal IV (1).
Appendix 2. Data matrix for cladistic analysis.
Coelophysis
?0100???0?10??????1?0000000?001011?00?00000001?0?00?000??
0?????00?1000100?00????0?001000111010?001000000?0000012?
?000001000???????00000120??00?000000000101010000000010?0
00001?000?00?0000000
Dilophosaurus
?1100??10?00?0????11000???0?10?001?00001?????000??01000??
??????000001000000001010?0001010010101011000000100000001
00000200?????????0000000000?0?00001000010?010000010010?0
00000000000000000000
Allosaurus
?1100?000000010001100010001011100111110010?000000101000
1000000000000101000100000000000010010101110001010100000
000000010000?01????0000000000010?00100001000001001001000
000?0100110000200000000
Sinraptor
?1100??00?000100010000000010?0100011110010?0?00000000001
0000?000000010000110?0000000000100101010100010101000000
000000????????2?00?0???00?0???????100001000?010?10?100000
0001000?0000200000000
Microraptor
0?????????????????10?0?????????????????????????????????????
?????0??010?0?0?1??????00??01100000????011??0?100001???01
10112111?01?101?011100?11010010001101111?0111031112100?2
111??0?0111101111
Sinornithosaurus
0011?????0??????????000???111010???10011110?1?0?001??????
??????00?0010000??1?00???00010110??100????????1?????0???0
0?1?1??????0111?11011000???0?001000?00101?0111103111210?
??1???????210001100
Velociraptor
?01101001000012011200000101110100111001211110?000010001
110100000000010000111100?000001011010100011001101111101
1100011011211100111011011000112000010001001111011100211
12100121?000000110100100
Shuuvuia
?01100000010??112011?000?0?0?200111?100?10001010001010??
?1?100101000100000002011000??10122??000???11111101???1?2
0 1 0 1 2 0 2 0/1 0 1 0 0 ? 2 0 0 0 ? 0 2 0 0 0 0 3 1 ? ? 2 1 2 2 1 1 0 1 1 0 0 0 0 ? 1 0 0 1 2 1 0 0 0 0 1 1
220??301001101311002000
Mononykus
?????00???????112???????????????????????????????????100????
?????????????????????????????2???0??????1??1101010??2?1??2?
???????2000?020000310?2122110110??0????1210?????2?0??3010
01101311002000
Patagonykus
????????????????????????????????????????????????????????????
????????????????????????????????????????????11??0??2?1??2??
????????????2000?310?210?1??110??0????1????????1000?201?00
0?0111100???
Confuciusornis
10110?????????????1??000?01??21001??0??2??0?0?10?0?01?????
??0??0000002000?10?0000001??1?????????0???1???01?002???0?
????2?111?21010??0111401121100100000111?1???110011122011
3?10??1?1301110010
Archaeopteryx
101?000??000??112010?010??111210?11?001210010??0?0??1001
11?1???000001010000020000000100120??0010011?1?0???0000??
?00110221000?1??1011011110?1200001000001111100?110311?11
01?2110000?01?0110010
Caudipteryx
00110?????????????0?0111??10?1?001?1???2100???00??????????
?????11110?200?????00???101?11???????0???01?0??????0???01?
???2?????0???0??0??0?0??200001000?00?1?????10?2??1??10?2??
???0??110000?00
Conchoraptor
?0110?????????1???001111?1???00111?00??210011000?0???1?0?
???10011110?20010?0?0001111??1???????????????01??10?2?110
??1???0??1???1?101?00000???????1?001001100011?0?????101??
2010?0??0??0?00??0
Citipati
?01101001001??2210001101111?000111?00102100110001010010
0011010011110?210100010001111??1?????????01101101111001?
?201??0210111?0111101001000?1200001000100111000??0?2??11
011?3010000001?0000000
Shenzhousaurus
???0??????????????2100001?10?0?0?1??000??000000???????????
?0????000001100??0?1??0?11??1?2???1?????????0???0?00???000
00?????????????????????????0??01000001??11?10?10000000?10
01?0????????????
Ornithomimus
?0101?110?101?10102100001010100001100000000000010100000
??????001000002000010?1110011??1???????0?0110?1001000?101
0000000101001?????021000200010?101020010000011010?100000
000100110010110001030
Gallimimus
?0101?1101101010102100001011?000?1100000000000010100000
1?010000100000200001021000011??1?????????01101100100000?
1000000010100??????0210002000?0?101010010000011010010000
0000100110010110001030
Troodon
???1?112?1101000001????0?011???0????20220002100?1??1100??
??0?1010??000?1??????????0???0111010100??1111011101?1?100
0?100?11????????????1?00???0?0?100010?????????0?2?11000?0
211000??01100011?1
Saurornithoides
?01???1??1101???0?1100001?10?0?0??????2????????????????1?0
10???10??000?10??1??????000001110101????????????0??1?100?
?1???????????????????????????????????????????0?2?01??0??211
0?????????01101
Sinovenator
?0???002?0000111100100101?1112?0011????2??011?0110?0100?
?????11000?000?1?????00?000000011110?1????1??100110000010
00110211?????????110111????2000???0????111?01111031111100
?2110000?01100001?1
Alxasaurus
????????????????????????????????????????????????????????????
???110?100?0?????????????????10?001?????1?0?100000?1?001??
2??1??????????0??000?110?00100011011?00?1?0?20112????1???
?00??????00020
Erlikosaurus
?0110?02?0?1?0??10100011100??00001?10000100010?0?1?0000?
?111100110010000000020000001??010100001??????????????????
???????????????????????0???????????????????????????????????
???????????0??0??
Albertosaurus
?1000?00?0???????1001000101?10000000110000?211?010010001
????0?00000010100?10001100000001001010111000?00010100000
10000001000?100010000000100010?001100010000010010?10010
0000100000010110001000
Tyrannosaurus
?1000?0110000100210010001010100000001100001211001001000
1?00000000000101001100011000000010010101110001000101000
001000000100??1?????000000100??0?10110001000001001001011
0000?100100010110001000
Ornitholestes
?0100??0?0?00?1???0?0010?01111?001?10001?0?1000001101011
????00000000101000?0?0000000110110101001???0110111000???
00??000?1??????????????010??????0?????1001?01??1011?011?0?
??????0??????000??0
Huaxiagnathus
?01?0?????????????00001???1??2??????00?????????????????????
????00?0010?0??????????00?001101010?0???????????01?????00?
?0?00010????000000000??10101100000010?0??01001000110??1?
????0??1100000?0
Sinosauropteryx
001?0?????????????00001???1??1??????0000?????0?1100???????
?????00??0101???????????001001101010?0??1?1?0???1010????0
0?10000010?????00000001??101111000?0010?????1001000110??
1?0??00??110000000
Compsognathus
? 0 1 ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 0 ? ? ? ? 1 1 1 ? ? 0 ? ? ? 0 0 0 0 0/1 1 ? ? 1 1 ? 0 ? ? ? 0 0 ? ? ? ? ?
??????0000010?0??10?0001?00100110101010??1?1?10??0010???0
00?0010001??????00?00??1??1010?1000?00???????10?1100010??
??1???0??210000000
Juravenator
? 0 1 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 0 0 ? ? ? 1 1 0 2 1 0 ? 1 ? ? ? ? 0 1/2 1 ? 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ?
? ? ? ? ? ? ? ? ? 0 ? 1 0 1 0 ? ? ? ? ? ? ? ? ? ? 0 0 0 ? 0 1 1 0 ? 0 1 ? ? 0 0 ? ? ? 1 ? ? ? ? ? ? 0 1 ? ? 0/1 0 0
0 0 ? 0 0 0 ? 0 0 0 0 ? ? ? ? ? 0 ? ? ? 0 0 2 0 ? ? ? 0 ? 0 1 1 0 0 0 0 0 0 0 0 ? 0 1 0 0/1 1 ? 0 ? ? ? ? ? ? ? ?
?101???0??????00000
Mirischia
????????????????????????????????????????????????????????????
????????????????????????????????????????????01?01???0??????
? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? 0 ? ? ? ? ? 0/1 1 0 1 1 ? 0 ? 0/1 1 0 0 ? 1 1 1 0
0??10?????????
Scipionyx
?0100?????????????0?001?????101001?0??00???0???0?1??1?????
??????0?0010000??0??0?0000??01?0??10??????1?????0??0??????
? 0 ? ? ? 0 0 0 ? ? ? ? ? 0 0 0 ? ? 0 1 0/1 ? ? ? 2 0 1 0 ? 1 0 0 0 0 ? ? 0 0 ? 0 ? ? ? 1 ? ? ? 0 ? 0 ? 1 0 ? ? ?
?????????????????
Nqwebasaurus
?0?????????????????????????????????????????????????????????
?????????????????????????????????????0???1?0???????????????
????????????00?00010?010?111010??????????????????200????0?
??0?11?000010
Coelurus
????????????????????????????????????????????????????????????
???????????????????????????????????????01000???????????????
?????????????????110????????????????????????????1000111010?
0?2100?????
Nedcolbertia
????????????????????????????????????????????????????????????
?????????????????????????????????????????????????????0?????
????????????????????????????0?????????????1???0?100?1??0000
001?0000??0
Supplementary Appendix 3. Character transformations for nodes of consensus tree discussed in Figure 4
of text (see Supplementary Appendix 1 for states):
Node 1: Root
Node 2:
char 2: 0 --> 1
char 11: 1 --> 0
char 29: 0 --> 1
char 33: 1 --> 0
char 67: 1 --> 0
char 69: 0 --> 1
char 85: 1 --> 0
char 88: 0 --> 1
char 89: 1 --> 0
char 90: 1 --> 0
char 97: 0 --> 1
char 111: 1 --> 0
char 112: 2 --> 0
char 136: 1 --> 0
char 137: 2 --> 0
char 163: 0 --> 1
char 174: 1 --> 0
Node 3:
char 19: 1 --> 0
char 27: 0 --> 1
char 75: 0 --> 1
char 101: 0 --> 1
char 146: 0 --> 1
char 153: 1 --> 0
char 160: 0 --> 1
char 172: 0 --> 1
char 181: 0 --> 1
Node 4:
char 36: 0 --> 1
char 103: 0 --> 1
char 181: 1 --> 2
Node 5:
char 2: 1 --> 0
char 89: 0 --> 1
char 97: 1 --> 0
char 99: 0 --> 1
Node 6:
char 31: 1 --> 0
char 95: 1 --> 0
char 122: 0 --> 1
char 132: 0 --> 2
char 158: 0 --> 1
Node 7 (Maniraptora):
char 36: 0 --> 1
char 40: 0 --> 1
char 51: 0 --> 1
char 53: 0 --> 1
char 104: 0 --> 1
char 106: 0 --> 1
char 166: 0 --> 1
Node 8:
char 4: 0 --> 1
char 45: 0 --> 1
char 71: 1 --> 0
char 93: 1 --> 0
char 119: 0 --> 2
char 157: 1 --> 0
char 165: 0 --> 1
Node 9:
char 9: 0 --> 1
char 123: 0 --> 1
char 124: 0 --> 1
char 127: 0 --> 1
char 132: 2 --> 1
char 153: 0 --> 1
Node 10
(Deinonychosauria):
char 28: 0 --> 1
char 31: 0 --> 1
char 39: 0 --> 1
char 57: 0 --> 1
char 121: 0 --> 1
char 131: 0 --> 1
char 134: 0 --> 1
char 136: 0 --> 1
char 156: 0 --> 1
char 161: 0 --> 1
char 168: 0 --> 1
char 173: 0 --> 1
Node 11
(Tyrannosauridae):
char 3: 1 --> 0
char 21: 0 --> 1
char 31: 1 --> 0
char 34: 1 --> 0
char 44: 0 --> 2
char 45: 0 --> 1
char 79: 0 --> 1
char 96: 0 --> 1
char 107: 0 --> 1
char 137: 0 --> 1
char 147: 0 --> 1
char 179: 0 --> 1
char 186: 0 --> 1
Node 12
(Compsognathidae):
char 28: 0 --> 1
char 30: 0 --> 2
char 109: 0 --> 1
char 145: 0 --> 1
Node 13
(Ornithomimosauria):
char 19: 0 --> 2
char 69: 1 --> 0
char 70: 0 --> 1
char 78: 0 --> 1
char 83: 0 --> 1
char 84: 0 --> 1
char 87: 0 --> 1
char 175: 0 --> 1
Node 14:
char 70: 1 --> 2
char 71: 1 --> 0
char 144: 0 --> 1
Node 15:
char 112: 1 --> 2
char 114: 0 --> 1
char 137: 0 --> 3
char 138: 0 --> 1
char 142: 0 --> 1
char 145: 0 --> 1
char 149: 0 --> 1
char 183: 0 --> 1
Node 16:
char 105: 1 --> 0
char 143: 0 --> 2
char 167: 1 --> 2
char 172: 2 --> 3
char 177: 0 --> 1
char 178: 0 --> 1
char 180: 0 --> 1
char 181: 1 --> 3
char 186: 0 --> 2
Node 17
(Oviraptorosauria):
char 22: 0 --> 1
char 24: 0 --> 1
char 64: 0 --> 1
char 66: 0 --> 1
char 67: 0 --> 1
char 70: 0 --> 2
char 83: 0 --> 1
char 87: 0 --> 1
char 115: 0 --> 1
char 169: 0 --> 1
char 161: 1 --> 0
char 163: 3 --> 2
Node 18:
char 21: 0 --> 1
char 32: 0 --> 1
char 33: 0 --> 1
char 36: 1 --> 0
char 84: 0 --> 1
char 110: 0 --> 1
char 130: 0 --> 1
Node 21:
char 58: 1 --> 0
char 76: 0 --> 1
char 85: 1 --> 0
char 187: 0 --> 1
Node 19 (Troodontidae):
char 8: 0 --> 2
char 20: 0 --> 1
char 62: 0 --> 1
char 69: 1 --> 0
char 72: 0 --> 1
char 90: 0 --> 1
char 94: 0 --> 1
Node 20:
char 7: 0 --> 1
char 10: 0 --> 1
char 11: 0 --> 1
char 13: 0 --> 1
char 17: 1 --> 0
char 64: 0 --> 1
char 91: 1 --> 0
char 92: 0 --> 1
char 110: 0 --> 1
Node 22
(Dromaeosauridae):
char 99: 1 --> 0
char 111: 0 --> 1
char 119: 2 --> 1
char 121: 1 --> 2
char 167: 1 --> 2
Node 23:
char 93: 0 --> 1
char 136: 1 --> 0
Node 24 (Aves):
char 1: 0 --> 1
char 30: 0 --> 2
char 45: 1 --> 0
char 120: 1 --> 2
char 152: 0 --> 1
char 170: 0 --> 1
char 185: 0 --> 1
char 188: 0 --> 1
Supplementary References. Below general references, there is a list of references for all taxa considered
in cladistic analysis
1. Weishampel, D. B., Dodson, P. & H. Osmólska eds. The Dinosauria. (Univ. California Press,
Berkeley, 2004). (2nd edition)
2. Nixon, K. 1999. The parsimony ratchet, a new method for rapid parsimony analysis.
Cladistics 15, 407-417.
3. Goloboff, P. 1993. NONA: A tree searching program. Program and documentation distributed
by the author, San Miguel de Tucumán, Argentina.
4. Goloboff, P. and J. Farris. Methods for quick consensus estimation. Cladistics 17, S26-S34
(2001).
5. Meyer, R. & Schmidt-Kaler, H. Paläogeographischer Atlas des süddeutschen Oberjura
(Malm). Geol. Jb. 115, 3-77 (1989).
Dilophosaurus wetherilli
6. Welles S. P. Dilophosaurus wetherilli (Dinosauria, Theropoda);osteology and comparisons.
Palaeontographica A185 (4-6):85-180 (1984).
Coelophysis bauri
7. Colbert E. H. The Triassic dinosaur Coelophysis. Bull. Mus. Northern Arizona 57, 160pp.
(1989).
Allosaurus fragilis
8. Madsen, J. H. Allosaurus fragilis: A revised osteology. Utah Dept. Natur. Survey Bull. 109, 1163 (1976).
Sinraptor dongi
9. Currie, P. J. & Zhao, X. A new carnosaur (Dinosauria, Theropoda) from the Jurassic of
Xinjiang, People's Republic of China. Can. J. Earth Sci. 30, 2037-2081 (1993).
Microraptor zhaoianus
10. Hwang, S. H., Norell, M. A., Qiang, J. & Keqin, G. New specimens of Microraptor zhaoianus
(Theropoda: Dromaeosauridae) from northeastern China. Am. Mus. Novitates 3381, 1-44
(2002).
Sinornithosaurus millenii
11. Xu X., Wang, X.-L. & Wu, X.-C. A dromaeosaurid dinosaur with a filamentous integument
from the Yixian Formation of China. Nature 401, 262-266 (1999).
Velociraptor mongoliensis
12. Norell, M. A. & Makovicky, P. J. Important features of the dromaeosaurid skeleton II:
Information from newly collected specimens of Velociraptor mongolensis. Am. Mus.
Novitates 3282, 1-45 (1999).
Shuuvuia deserti
13. Chiappe, L. M., Norell, M. A. & Clark, J. M. The skull of a new relative of the stem-group
bird Mononykus. Nature 392, 275-278 (1998).
14. Suzuki, S. et al. A new specimen of Shuvuuia desteri Chiappe et al., 1998 from the Mongolian
Late Cretaceous with a discussion of the relationships of Alvarezsaurids to other theropod
dinosaurs. Contrib. Sci Natur. Hist. Mus. Los Angeles County 494, 1-18 (2002).
15. Schweitzer, M. H. et al. Beta-keratin specific immunological reactivity in feather-like
structures of the Cretaceous alvarezsaurid, Shuvuuia deserti. J. Exp. Zool. 285, 146-157
(1999).
Mononykus olecranus
16. Perle, A, Chiappe, L. M., Barsbold, R., Clark, J. M.& Norell, M. A. Skeletal morphology of
Mononykus olecranus (Theropoda: Aviale) from the Late Cretaceous of Mongolia. Am Mus
Novitates. 3105, 1-29 (1994).
17. Perle, A, Norell, M. A., Chiappe, L. M. & Clark, J. M. Flightless bird from the Cretaceous of
Mongolia. Nature 362, 623-626 (1993).
Patagonykus puertai
18 Chiappe, L. M. & Coria, R. A. A new speciemen of Patagonykus puertai (Theropoda:
Alvarezsauridae) from the Late Cretaceous of Patagonia. Ameghiniana 40, 119-122 (2003).
Confuciusornis sanctus
19. Chiappe, L. M., Ji, S., Ji, Q. & Norell, M. A. Anatomy and systematics of the
Confuciornithidae (Theropoda: Aves) from the Late Mesozoic of northeastern China. Bull.
Am. Mus. Natur. Hist. 242, 1-89 (1999).
Archeopteryx lithographica
20. Wellnhofer, P. Das fünfte Skelettexemplar von Archaeopteryx. Palaeontographica (A) 147,
169-216 (1974).
21. Wellnhofer, P. A New Specimen of Archaeopteryx. Science 240, 1790-1792 (1988).
22. Wellnhofer, P. Ein neues Exemplar von Archaeopteryx. Archaeopteryx 6, 1-30 (1988).
23. Wellnhofer, P. in Papers in Avian Paleontology (ed Campbell, Jr., K. E.), 3-23, Natur. Hist.
Mus. Los Angeles County, Sci. Ser. 36 (1992).
24. Wellnhofer, P. Das siebte Exemplar von Archaeopteryx aus den Solnhofener Schichten.
Archaeopteryx 11, 1-48 (1993).
25. Elzanowski, A. & Wellnhofer, P. Cranial morphology of Archaeopteryx: Evidence from the
seventh specimen. J. Vertebr. Paleont. 16(1), 81-94 (1996).
Caudipteryx zoui
26. Ji, Q., Currie, P. J., Norell, M. A. & Ji S.-A. Two feathered theropods from the Upper
Jurassic/Lower Cretaceous strata of northeastern China. Nature 393, 753-761 (1998).
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